Mimicry occurs when a group of organisms,[3] the mimics, evolve to share common perceived characteristics with another group, the models.[4] The evolution is driven by the selective action of a signal-receiver or dupe.[5] Birds, for example, use sight to identify palatable insects (the mimics), whilst avoiding the noxious models.

The model is usually another species except in cases of automimicry. The deceived signal-receiver is typically another organism, such as the common predator of two species.[a] As an interaction, mimicry is in most cases advantageous to the mimic and harmful to the receiver, but may increase, reduce or have no effect on the fitness of the model depending on the situation. Models themselves can be difficult to define; for example, eye spots may not bear resemblance to any specific organism's eyes, and camouflage often cannot be attributed to a particular model.

Mimicry is related to camouflage, in which a species resembles its surroundings or is otherwise difficult to detect. In particular, mimesis, in which the mimic takes on the properties of a specific object or organism, but one to which the dupe is indifferent, is an area of overlap between camouflage and mimicry.[7] For example, animals such as flower mantises, planthoppers and geometer moth caterpillars that resemble twigs, bark, leaves or flowers are mimetic.[2]p51 The difficulty is sometimes avoided by choosing a different term; "crypsis" (in the broad sense) is sometimes used to encompass all forms of avoiding detection, such as mimicry, camouflage and hiding.[8][8]

Mimics may have different models for different life cycle stages, or they may be polymorphic, with different individuals imitating different models. Models themselves may have more than one mimic, though frequency dependent selection favors mimicry where models outnumber mimics. Models tend to be relatively closely related organisms,[9] but mimicry of vastly different species is also known. Most known mimics are insects,[2] though many other animal mimics, including mammals, are known. Plants and fungi may also be mimics, though less research has been carried out in this area.[10][11][12]

Use of the word mimicry dates to 1637. It derives from the Greek term mimetikos, "imitative", in turn from mimetos, the verbal adjective of mimeisthai, "to imitate". Originally used to describe people, "mimetic" was used in zoology from 1851, "mimicry" from 1861.[13]

Many types of mimicry have been described. An overview of each follows, highlighting the similarities and differences between the various forms. Classification is often based on function with respect to the mimic (e.g., avoiding harm). Some cases may belong to more than one class, e.g., automimicry and aggressive mimicry are not mutually exclusive, as one describes the species relationship between model and mimic, while the other describes the function for the mimic (obtaining food).

Defensive or protective mimicry takes place when organisms are able to avoid harmful encounters by deceiving enemies into treating them as something else. The first three such cases discussed here entail mimicry of organisms protected by warning coloration: Batesian mimicry, where a harmless mimic poses as harmful; Müllerian mimicry, where two or more harmful species mutually advertise themselves as harmful; and Mertensian mimicry, where a deadly mimic resembles a less harmful but lesson-teaching model. The fourth case, Vavilovian mimicry, where weeds resemble crops, is important for several reasons; and humans are the agent of selection.

In Batesian mimicry the mimic shares signals similar to the model, but does not have the attribute that makes it unprofitable to predators (e.g., unpalatability). In other words, a Batesian mimic is a sheep in wolf's clothing. It is named after Henry Walter Bates, an English naturalist whose work on butterflies in the Amazon rainforest (including Naturalist on the River Amazons) was pioneering in this field of study.[15][16] Mimics are less likely to be found out when in low proportion to their model, a phenomenon called negative frequency dependent selection that applies in most other forms of mimicry as well. Batesian mimicry can only be maintained if the harm caused to the predator by eating a model outweighs the benefit of eating a mimic. The nature of learning is weighted in favor of the mimics, for a predator that has a bad first experience with a model tends to avoid anything that looks like it for a long time and does not re-sample soon to see whether the initial experience was a false negative. However, if mimics become more abundant than models, then the probability of a young predator having a first experience with a mimic increases. Such systems are therefore most likely to be stable where both the model and the mimic occur and where the model is more abundant than the mimic.[17] This is not the case in Müllerian mimicry however, which is described next. There are many Batesian mimics in the Lepidoptera. Consul fabius and Eresia eunice imitate unpalatable Heliconius butterflies such as H. ismenius.[18] Several palatable moths produce ultrasonic click calls to mimic unpalatable tiger moths.[19]Octopuses of the genus Thaumoctopus (the mimic octopus) are able to intentionally alter their body shape and coloration to resemble dangerous sea snakes or lionfish.[20]

Müllerian mimicry describes a situation where two or more species have similar warning or aposematic signals and both share genuine anti-predation attributes (e.g., being unpalatable). At first, Bates could not explain why this should be so—if both were harmful why did one need to mimic another? The German naturalist Fritz Müller put forward the first explanation for this phenomenon: If a common predator confuses two species, individuals in both those speces are more likely to survive.[22][23] This type of mimicry is unique in several respects. Firstly, both the mimic and the model benefit from the interaction, which could thus be classified as mutualism in this respect. The signal receiver is also advantaged by this system, despite being deceived regarding species identity, as it avoids potentially harmful encounters. The usually clear identity of mimic and model are also blurred. In cases where one species is scarce and another abundant, the rare species can be said to be the mimic. When both are present in similar numbers however it is more realistic to speak of each as comimics than of a distinct 'mimic' and 'model' species, as their warning signals tend to converge toward something intermediate between the two.[24] Also, the two species may exist on a continuum from harmless to highly noxious, so Batesian mimicry grades smoothly into Müllerian convergence.[25][26]

The monarch butterfly (Danaus plexippus) is a member of a Müllerian complex with the viceroy butterfly (Limenitis archippus) in shared coloration patterns and display behavior. The viceroy has subspecies with somewhat different coloration, each one very closely matching the local Danaus species. E.g., in Florida, the pairing is of the viceroy and the queen butterfly, and in Mexico, the viceroy resembles the soldier butterfly. Therefore, the viceroy is a single species involved in three different Müllerian pairs.[27] This example was long believed to be Batesian, with the viceroy mimicking the monarch, but the viceroy is actually the more unpalatable species.[28] The genus Morpho is palatable, but some species (such as M. amathonte) are very strong fliers; birds – even species that specialize in catching butterflies on the wing – find it very hard to catch them.[29] The conspicuous blue coloration shared by most Morpho species may be Müllerian, [18] or may be "pursuit aposematism".[30] The "orange complex" of distasteful butterfly species includes the heliconiinesAgraulis vanillae, Dryadula phaetusa, and Dryas iulia.[18] At least seven species of millipedes in the genera Apheloria and Brachoria (Xystodesmidae) form a Müllerian mimicry ring in the eastern United States, in which unrelated polymorphic species converge on similar color patterns where their range overlaps.[31]

Emsleyan[7] or Mertensian mimicry describes unusual cases where deadly prey mimic a less dangerous species. It was first proposed by Emsley[32] as a possible answer for the theoretical difficulties a predator species faces when associating an aposematic phenotype of potentially dangerous animals, such as the coral snake, with unprofitability when the predator has an increased risk of death, negating any learned behaviour. The theory was developed by the German biologist Wolfgang Wickler in a chapter of Mimicry in Plants and Animals,[2] who named it after the GermanherpetologistRobert Mertens.[33] Sheppard points out that Hecht and Marien put forward a similar hypothesis ten years earlier.[34][35] This scenario is a little more difficult to understand, as in other types of mimicry it is usually the most harmful species that is the model. But if a predator dies, it cannot learn to recognize a warning signal, e.g., bright colors in a certain pattern. In other words, there is no advantage in being aposematic for an organism that is likely to kill any predator it succeeds in poisoning; such an animal is better off being camouflaged, to avoid attacks altogether. If, however, there were some other species that is harmful but not deadly as well as aposematic, the predator can learn to recognize its particular warning colors and avoid such animals. A deadly species can then profit by mimicking the less dangerous aposematic organism if this reduces the number of attacks.[34][35] The exception here, ignoring any chance of animals learning by watching a conspecific die (see Jouventin et al. for a discussion of observational learning and mimicry),[36] is the possibility of not having to learn that it is harmful in the first place: instinctive genetic programming to be wary of certain signals. In this case, other organisms could benefit from this programming, and Batesian or Müllerian mimics of it could potentially evolve. In fact, it has been shown that some species do have an innate recognition of certain aposematic warnings. Hand-reared turquoise-browed motmots (Eumomota superciliosa), avian predators, instinctively avoid snakes with red and yellow rings.[37] Other colors with the same pattern, and even red and yellow stripes with the same width as rings, were tolerated. However, models with red and yellow rings were feared, with the birds flying away and giving alarm calls in some cases. This provides an alternative explanation to Mertensian mimicry. See Greene and McDiarmid for a review of the subject.[38]

Some harmless milk snake (Lampropeltis triangulum) subspecies, the moderately toxic false coral snakes (genus Erythrolamprus), and the deadly coral snakes (genus Micrurus) all have a red background color with black and white / yellow rings. In this system, both the milk snakes and the deadly coral snakes are mimics, whereas the false coral snakes are the model.[32]

Wasmannian mimicry refers to cases where the mimic resembles a model along with which it lives (inquiline) in a nest or colony. Most of the models here are social insects such as ants, termites, bees and wasps.[39]

Vavilovian mimicry presents an illustration of unintentional (or rather 'anti-intentional') selection by man. While some cases of artificial selection go in the direction desired, such as selective breeding, this case presents the opposite characteristics. Weeders do not want to select weeds that look increasingly like the cultivated plant, yet there is no other option. One case is Echinochloa oryzoides, a species of grass found as a weed in rice (Oryza sativa) fields. The plant looks similar to rice; its seeds are often mixed in rice and have become difficult to separate through Vavilovian mimicry.[41] A similar problem in agriculture is pesticide resistance: farmers do not wish to select for weeds that have increasingly similar resistance to pesticides as the crop itself, yet that is the inevitable effect. Vavilovian mimics may eventually be domesticated themselves, as in the case of rye in wheat; Vavilov called these weed-crops secondary crops.

Vavilovian mimicry can be classified as defensive mimicry, in that the weed mimics a protected species. This bears strong similarity to Batesian mimicry in that the weed does not share the properties that give the model its protection, and both the model and the dupe (in this case people) are harmed by its presence. There are some key differences, though; in Batesian mimicry the model and signal receiver are enemies (the predator would eat the protected species if it could), whereas here the crop and its human growers are in a mutualistic relationship: the crop benefits from being dispersed and protected by people, despite being eaten by them. In fact, the crop's only "protection" relevant here is its usefulness to humans. Secondly, the weed is not eaten, but simply destroyed. The only motivation for killing the weed is its effect on crop yields. Finally, this type of mimicry does not occur in ecosystems unaltered by humans.

Unlike the previously mentioned forms of mimicry, Gilbertian mimicry involves only two species. The potential host / prey drives away its parasite / predator by mimicking it, the reverse of host-parasite aggressive mimicry. It was coined by Pasteur as a phrase for such rare mimicry systems,[7] and is named after the American ecologist Lawrence E. Gilbert.[42]

This form of protective mimicry occurs in the genus Passiflora. The leaves of this plant contain toxins that deter herbivorous animals—however some Heliconius butterfly larvae have evolved enzymes that break down these toxins, allowing them to specialize on this genus. This has created further selection pressure on the host plants, which have evolved stipules that mimic mature Heliconius eggs near the point of hatching. These butterflies tend to avoid laying eggs near each existing ones, which helps avoid exploitative intraspecific competition between caterpillars — those that lay on vacant leaves provide their offspring with a greater chance of survival. Most Heliconius larvae are cannibalistic, meaning that on leaves older eggs hatch first and eat the new arrivals. Thus, it seems that such plants have evolved egg dummies due to these grazing herbivore enemies. In addition, the decoy eggs are also nectaries, attracting predators of the caterpillars such as ants and wasps.[9] This acts as a further defense of the plant against the caterpillars.

Browerian mimicry,[7] named after Lincoln P. Brower and Jane Van Zandt Brower,[43][44] is a form of automimicry; where the model belongs to the same species as the mimic. This is the analogue of Batesian mimicry within a single species, and occurs when there is a palatability spectrum within a population. Examples include the monarch and the queen from the Danainae subfamily, which feed on milkweed species of varying toxicity. These species store toxins from its host plant, which are maintained even in the adult (imago) form. As levels of toxin vary depending on diet during the larval stage, some individuals are more toxic than others. Less palatable organisms, therefore, mimic more dangerous individuals, with their likeness already perfected.

This is not always the case, however. In sexually dimorphic species, one sex may be more of a threat than the other, which could mimic the protected sex. Evidence for this possibility is provided by the behavior of a monkey from Gabon, which regularly ate male moths of the genus Anaphe, but promptly stopped after it tasted a noxious female.[45]

Aggressive mimicry describes predators (or parasites) that share the same characteristics as a harmless species, allowing them to avoid detection by their prey (or host). The mimic may resemble the prey or host itself, or another organism that is either neutral or beneficial to the signal receiver. In this class of mimicry the model may be affected negatively, positively or not at all. Just as parasites can be treated as a form of predator,[46] host-parasite mimicry is treated here as a subclass of aggressive mimicry.

The mimic may have a particular significance for duped prey. One such case is spiders, amongst which aggressive mimicry is quite common in both luring prey and disguising stealthily approaching predators.[47] One case is the golden orb weaver (Nephila clavipes), which spins a conspicuous golden colored web in well-lit areas. Experiments show that bees are able to associate the webs with danger when the yellow pigment is not present, as occurs in less well-lit areas where the web is much harder to see. Other colors were also learned and avoided, but bees seemed least able to effectively associate yellow pigmented webs with danger. Yellow is the color of many nectar bearing flowers, however, so perhaps avoiding yellow is not worthwhile. Another form of mimicry is based not on color but pattern. Species such as Argiope argentata employ prominent patterns in the middle of their webs, such as zigzags. These may reflect ultraviolet light, and mimic the pattern seen in many flowers known as nectar guides. Spiders change their web day to day, which can be explained by bee's ability to remember web patterns. Bees are able to associate a certain pattern with a spatial location, meaning the spider must spin a new pattern regularly or suffer diminishing prey capture.[48]

Another case is where males are lured towards what seems to be a sexually receptive female. The model in this situation is the same species as the dupe. Beginning in the 1960s, James E. Lloyd's investigation of female fireflies of the genus Photuris revealed they emit the same light signals that females of the genus Photinus use as a mating signal.[49] Further research showed male fireflies from several different genera are attracted to these "femmes fatales", and are subsequently captured and eaten. Female signals are based on that received from the male, each female having a repertoire of signals matching the delay and duration of the female of the corresponding species. This mimicry may have evolved from non-mating signals that have become modified for predation.[50]

The spotted predatory katydid (Chlorobalius leucoviridis) is an acoustic aggressive mimic of cicadas.

The listrosceline katydidChlorobalius leucoviridis of inland Australia is capable of attracting male cicadas of the tribe Cicadettini by imitating the species-specific reply clicks of sexually receptive female cicadas. This example of acoustic aggressive mimicry is similar to the Photuris firefly case in that the predator's mimicry is remarkably versatile – playback experiments show that C. leucoviridis is able to attract males of many cicada species, including Cicadettine cicadas from other continents, even though cicada mating signals are species-specific.[51]

Luring is not a necessary condition however, as the predator still has a significant advantage simply by not being identified as such. They may resemble a mutualistic symbiont or a species of little relevance to the prey.

A case of the latter situation is a species of cleaner fish and its mimic, though in this example the model is greatly disadvantaged by the presence of the mimic. Cleaner fish are the allies of many other species, which allow them to eat their parasites and dead skin. Some allow the cleaner to venture inside their body to hunt these parasites. However, one species of cleaner, the bluestreak cleaner wrasse (Labroides dimidiatus), is the unknowing model of a mimetic species, the sabre-toothed blenny (Aspidontus taeniatus). This wrasse, shown to the right cleaning a grouper of the genus Epinephelus, resides in coral reefs in the Indian and the Pacific Oceans, and is recognized by other fishes that then let it clean them. Its imposter, a species of blenny, lives in the Indian Ocean—and not only looks like it in terms of size and coloration, but even mimics the cleaner's "dance". Having fooled its prey into letting its guard down, it then bites it, tearing off a piece of its fin before fleeing. Fish grazed on in this fashion soon learn to distinguish mimic from model, but because the similarity is close between the two they become much more cautious of the model as well, so both are affected. Due to victims' ability to discriminate between foe and helper, the blennies have evolved close similarity, right down to the regional level.[53]

Another interesting example that does not involve any luring is the zone-tailed hawk, which resembles the turkey vulture. It flies amongst the vultures, suddenly breaking from the formation and ambushing its prey.[54] Here the hawk's presence is of no evident significance to the vultures, affecting them neither negatively or positively.

Parasites can also be aggressive mimics, though the situation is somewhat different from those outlined previously.

Some of the predators described have a feature that draws prey, and parasites can also mimic their host's natural prey, but are eaten themselves, a pathway into their host. Leucochloridium, a genus of flatworm, matures in the digestive system of songbirds, their eggs then passing out of the bird via the feces. They are then taken up by Succinea, a terrestrial snail. The eggs develop in this intermediate host, and then must find of a suitable bird to mature in. As the host birds do not eat snails, so the sporocyst has another strategy to reach its host's intestine. They are brightly colored and move in a pulsating fashion. A sporocyst-sac pulsates in the snail's eye stalks,[55][56] coming to resemble an irresistible meal for a songbird. In this way, it can bridge the gap between hosts, allowing it to complete its life cycle.[2] A nematode (Myrmeconema neotropicum) changes the colour of the abdomen of workers of the canopy ant Cephalotes atratus to make it appear like the ripe fruits of Hyeronima alchorneoides. It also changes the behaviour of the ant so that the gaster (rear part) is held raised. This presumably increases the chances of the ant being eaten by birds. The droppings of birds are collected by other ants and fed to their brood, thereby helping to spread the nematode.[57]

In an unusual case, planidium larvae of some beetles of the genus Meloe form a group and produce a pheromone that mimics the sex attractant of its host bee species. When a male bee arrives and attempts to mate with the mass of larvae, they climb onto his abdomen. From there, they transfer to a female bee, and from there to the bee nest to parasitize the bee larvae.[58]

Host-parasite mimicry is a two species system where a parasite mimics its own host. Cuckoos are a canonical example of brood parasitism, a form of kleptoparasitism where the mother has its offspring raised by another unwitting organism, cutting down the biological mother's parental investment in the process. The ability to lay eggs that mimic the host eggs is the key adaptation. The adaptation to different hosts is inherited through the female line in so-called gentes. Cases of intraspecific brood parasitism, where a female lays in conspecific's nest, as illustrated by the goldeneye duck (Bucephala clangula),[59] do not represent a case of mimicry. Another example is that of chemical mimicry, in which the parasitic butterfly Phengaris rebeli, which parasitizes the ant species Myrmica schencki by releasing chemicals that fool the worker ants to believe that the caterpillar larvae are ant larvae, and enable the P. rebeli larvae to be brought directly into the M. schencki nest.[60] Parasitic (cuckoo) bumblebees (formerly Psithyrus, now included in Bombus) resemble their hosts more closely than would be expected by chance, at least in areas like Europe where parasite-host co-speciation is common; but this is explainable as Müllerian mimicry, rather than requiring the parasite's coloration to deceive the host and thus constitute aggressive mimicry.[61]

Reproductive mimicry occurs when the actions of the dupe directly aid in the mimic's reproduction. This is common in plants with deceptive flowers that do not provide the reward they seem to. Other forms of mimicry have a reproductive component, such as Vavilovian mimicry involving seeds, and brood parasitism, which also involves aggressive mimicry.[citation needed]

Bakerian mimicry, named after Herbert G. Baker,[62] is a form of automimicry where female flowers mimic male flowers of their own species, cheating pollinators out of a reward. This reproductive mimicry may not be readily apparent as members of the same species may still exhibit some degree of sexual dimorphism. It is common in many species of Caricaceae.[63]

Like Bakerian mimicry, Dodsonian mimicry is a form of reproductive floral mimicry, but the model belongs to a different species than the mimic. The name refers to Calaway H. Dodson.[64] By providing similar sensory signals as the model flower, it can lure its pollinators. Like Bakerian mimics, no nectar is provided. Epidendrum ibaguense of the family Orchidaceae resembles flowers of Lantana camara and Asclepias curassavica, and is pollinated by monarch butterflies and perhaps hummingbirds.[65] Similar cases are seen in some other species of the same family. The mimetic species may still have pollinators of its own though. For example a lamellicorn beetle, which usually pollinates correspondingly colored Cistus flowers, is also known to aid in pollination of Ophrys species that are normally pollinated by bees.[66]

Pseudocopulation occurs when a flower mimics a female of a certain insect species, inducing the males to try to copulate with the flower. This is much like the aggressive mimicry in fireflies described previously, but with a more benign outcome for the pollinator. This form of mimicry has been called Pouyannian mimicry,[7] after Maurice-Alexandre Pouyanne, who first described the phenomenon.[67][68] It is most common in orchids, which mimic females of the order Hymenoptera (generally bees and wasps), and may account for around 60% of pollinations.[69] Depending on the morphology of the flower, a pollen sac called a pollinia is attached to the head or abdomen of the male. This is then transferred to the stigma of the next flower the male tries to inseminate, resulting in pollination. Visual mimicry is the most obvious sign of this deception for humans, but the visual aspect may be minor or non-existent. It is the senses of touch and olfaction that are most important.[69]

Inter-sexual mimicry occurs when individuals of one sex in a species mimic members of the opposite sex. An example is the three male forms of the marine isopodParacerceis sculpta. Alpha males are the largest and guard a harem of females. Beta males mimic females and manage to enter the harem of females without being detected by the alpha males allowing them to mate. Gamma males are the smallest males and mimic juveniles. This also allows them to mate with the females without the alpha males detecting them.[70] Similarly, among common side-blotched lizards, some males mimic the yellow throat coloration and even mating rejection behavior of the other sex to sneak matings with guarded females. These males look and behave like unreceptive females. This strategy is effective against "usurper" males with orange throats, but ineffective against blue throated "guarder" males, which chase them away.[71][72] Female spotted hyenas have pseudo-penises that make them look like males.[73]

Automimicry: gray hairstreak (Strymon melinus), showing the false head at the rear, held upwards at rest

Effective deflection of attacks: purple hairstreak, Neozephyrus quercus, showing damage from a bird attack at the wrong end

Automimicry or intraspecific mimicry occurs within a single species. One form of such mimicry is where one part of an organism's body resembles another part. For example, the tails of some snakes resemble their heads and they show behavior such as moving backwards when threatened and presenting the predator with the tail, thereby improving their chances of escape without fatal harm. Some species of fishes have eyespots near their tails, and when mildly alarmed swim slowly backwards, presenting the tail as a head. Some insects too, have tail patterns and appendages of various degrees of sophistication that promote attacks at the rear rather than at the head.

Some writers use the term "automimicry" when the mimic imitates other morphs within the same species. For example, in a species where males mimic females or vice versa, this may be an instance of sexual mimicry in evolutionary game theory. Examples are found in some species of birds, fishes, and lizards.[74] Quite elaborate strategies along these lines are known, such as in the notorious "scissors, paper, rock" example in the species Uta stansburiana,[75] but there are qualitatively different examples in many other species, such as some Platysaurus.[76]

Many species of insects are toxic or distasteful when they have fed on certain plants that contain chemicals of particular classes, but not when they have fed on plants that lack those chemicals. For instance, some species of the subfamilyDanainae feed on various species of the Asclepiadoideae in the family Apocynaceae, which render them poisonous and emetic to most predators. Such insects frequently are aposematically coloured and patterned. When feeding on innocuous plants however, they are harmless and nutritious, but a bird that once has sampled a toxic specimen is unlikely to eat harmless specimens that have the same aposematic coloration. When regarded as mimicry of toxic members of the same species, this too may be seen as automimicry.[77]

For example, several species of pygmy owls bear "false eyes" on the back of the head, misleading predators into reacting as though they were the subject of an aggressive stare.[citation needed] Some species of caterpillar, such as many of the hawkmoths (Sphingidae), have eyespots on their anterior abdominal segments. When alarmed, they retract the head and the thoracic segments into the body, leaving the apparently threatening large eyes at the front of the visible part of the body.[citation needed] Many insects have filamentous "tails" at the ends of their wings and patterns of markings on the wings themselves. These combine to create a "false head". This misdirects predators such as birds and jumping spiders (Salticidae). Spectacular examples occur in the hairstreak butterflies; when perching on a twig or flower, they commonly do so upside down and shift their rear wings repeatedly, causing antenna-like movements of the "tails" on their wings. Studies of rear-wing damage support the hypothesis that this strategy is effective in deflecting attacks from the insect's head.[78][79]

Some forms of mimicry do not fit easily within the classification given previously.

Floral mimicry is induced by the discomycete fungusMonilinia vaccinii-corymbosi.[80] In this unusual case, a fungal plant pathogen infects leaves of blueberries, causing them to secrete sugars, in effect mimicking the nectar of flowers. To the naked eye the leaves do not look like flowers, yet they still attract pollinating insects like bees using an ultraviolet signal. This case is unusual, in that the fungus benefits from the deception but it is the leaves that act as mimics, being harmed in the process. It is similar to host-parasite mimicry, but the host does not receive the signal. It has a little in common with automimicry, but the plant does not benefit from the mimicry, and the action of the pathogen is required to produce it.[80]

It is widely accepted that mimicry evolves as a positive adaptation. The lepidopterist and novelistVladimir Nabokov argued that although natural selection might stabilize a "mimic" form, it would not be necessary to create it.[81]

The most widely accepted model used to explain the evolution of mimicry in butterflies is the two-step hypothesis. The first step involves mutation in modifier genes that regulate a complex cluster of linked genes that cause large changes in morphology. The second step consists of selections on genes with smaller phenotypic effects, creating an increasingly close resemblance. This model is supported by empirical evidence that suggests that a few single point mutations cause large phenotypic effects, while numerous others produce smaller effects. Some regulatory elements collaborate to form a supergene for the development of butterfly color patterns. The model is supported by computational simulations of population genetics.[82] The Batesian mimicry in Papilio polytes is controlled by the doublesex gene.[83]

Some mimicry is imperfect. Natural selection drives mimicry only far enough to deceive predators. For example, when predators avoid a mimic that imperfectly resembles a coral snake, the mimic is sufficiently protected.[84][85][86]

^M. Edmunds. 1974. Defence in Animals: a survey of anti-predator defences. Harlow, Essex and New York, Longman. ISBN 0-582-44132-3. On p 255–256 there is a discussion of "pursuit aposematism": "Young suggested that the brilliant blue colours and bobbing flight of Morpho butterflies may induce pursuit... Morpho amathonte is a very fast flier... It is possible that birds that have chased several unsuccessfully may learn not to pursue butterflies of that [type]... In one area, Young found that 80% of less brilliant species of Morpho had beak marks on their wings... but none out of 31 M. amathonte... "If brilliant colour was a factor in courtship, then the conflicting selection pressures of sexual selection and predator selection might lead to different results in quite closely related species".